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New Eocene Primate from Myanmar Shares Dental Characters with African Eocene Crown Anthropoids

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New Eocene Primate from Myanmar Shares Dental Characters with African Eocene Crown Anthropoids ARTICLE https://doi.org/10.1038/s41467-019-11295-6 OPEN New Eocene primate from Myanmar shares dental characters with African Eocene crown anthropoids Jean-Jacques Jaeger1, Olivier Chavasseau 1, Vincent Lazzari1, Aung Naing Soe2, Chit Sein 3, Anne Le Maître 1,4, Hla Shwe5 & Yaowalak Chaimanee1 Recent discoveries of older and phylogenetically more primitive basal anthropoids in China and Myanmar, the eosimiiforms, support the hypothesis that Asia was the place of origins of 1234567890():,; anthropoids, rather than Africa. Similar taxa of eosimiiforms have been discovered in the late middle Eocene of Myanmar and North Africa, reflecting a colonization event that occurred during the middle Eocene. However, these eosimiiforms were probably not the closest ancestors of the African crown anthropoids. Here we describe a new primate from the middle Eocene of Myanmar that documents a new clade of Asian anthropoids. It possesses several dental characters found only among the African crown anthropoids and their nearest rela- tives, indicating that several of these characters have appeared within Asian clades before being recorded in Africa. This reinforces the hypothesis that the African colonization of anthropoids was the result of several dispersal events, and that it involved more derived taxa than eosimiiforms. 1 Laboratory PALEVOPRIM, UMR CNRS 7262, University of Poitiers, 6 rue Michel Brunet Cedex 9, 86073 Poitiers, France. 2 University of Distance Education, Mandalay 05023, Myanmar. 3 Ministry of Education, Department of Higher Education, Naypyitaw 15011, Myanmar. 4 Department of Theoretical Biology, University of Vienna, Althanstrasse 14, 1090 Vienna, Austria. 5 Department of Archaeology and National Museum, Mandalay Branch, Ministry of Religious Affairs and Culture, Mandalay 05011, Myanmar. Correspondence and requests for materials should be addressed to J.-J.J. (email: [email protected]) NATURE COMMUNICATIONS | (2019) 10:3531 | https://doi.org/10.1038/s41467-019-11295-6 | www.nature.com/naturecommunications 1 ARTICLE NATURE COMMUNICATIONS | https://doi.org/10.1038/s41467-019-11295-6 he evolutionary history of anthropoids, in particular their Diagnosis Medium-sized anthropoid (similar in tooth size to geographical origin, has been intensely debated during the Ateles with estimated body weight between 2.8 and 3.4 kg) T 1–5 last decades . Earlier hypotheses advocating an African (Supplementary Note 1) characterized by its short and elevated origin for anthropoids have been overturned by the discovery of muzzle, small and forwardly located orbits with low convergence older and phylogenetically more basal anthropoids in Asia, which (Supplementary Note 2), large upper canine with mesial groove, are thought to comprise the clade Eosimiiformes1,3,6–8. Recent three premolars with reduced unicuspid P2,P3, and P4 unwaisted, findings in Libya and Myanmar3,9 have shown that the eosimii- buccolingually broad with a small, mesially located protocone, no forms dispersed from Asia to Africa during the Middle Eocene. lingual cingulum, and reduced styles and buccal cingula. Anthropoids are also represented in the Southeast Asian Eocene Subrectangular M2 with large trigon basin, peripheralized and by another group, the amphipithecids10–14, the affinities of low cusps, U-shaped protocristae, complete and low crests which remain a matter of debate. However, crown anthropoids without paraconule but with a tiny swelling, which may and their nearest fossil relatives do not appear to be closely correspond to a vestigial metaconule, reduced buccal cingulum, related to these eosimiiforms, leaving their ancestors unknown. strong and continuous mesial, lingual and distal cingulum They appear successively in the African fossil record without hypocone or pericone, and weak hypoparacrista. Upper (proteopithecids, parapithecoids, oligopithecids, and proplio- molars trigone basins with slight enamel crenulations. Deep lower pithecids) between the late middle Eocene and the earliest Oli- jaw with most probably unfused symphysis, large canine alveolus, gocene. Available data converge to support multiple dispersal P2-P3 with strong protoconids surrounded by complete cingulids, events from Asia to Africa by anthropoid taxa, which were more stronger lingually than buccally, and with small paraconids and derived than eosimiiforms. hypoconids. P2 with drop-like occlusal outline, massive single 2 A new primate was collected from the Pondaung Formation, root with crown surface similar to that of P3.P2 larger than P .P3 about 2 km thick continental formation of Central Myanmar with stronger hypoconid, without talonid basin nor metaconid. from which only lower half of the Upper Member has yielded M3 with very short trigonid basin and large and deep talonid fossil mammals15. These fossiliferous deposits have been recently basin, with a large, central hypoconulid. Entoconid fused into radiometrically redated from 40.31 to 40.22 Ma (late Middle entocristid, closing the talonid lingually. Main cusps bunodont Eocene)16 and are therefore older than any known African with rounded wear facets. anthropoid locality. The new fossils were found in Paukkaung It differs from the eosimiiforms by its larger size, larger P2,P3 Kyitchaung 2 (PK2) locality, nearby the village of Paukkaung with a more extended lingual lobe and a distinct protocone, upper (Myaing township, Magway Region), a rich mammalian locality molars with small styles and buccal cingula, less distinct that has previously yielded several remains of other anthropoid hypoparacrista, more bunodont and peripheralized cusps, sub- primates such as eosimiiforms, amphipithecines, and also siva- rectangular M2 outline. It differs from the amphipithecines by its ladapid strepsirrhines3,12,17–19. They have been excavated from a lower premolar structure and proportions and by the absence of a fluviatile clayish and sandy layer and were associated with diverse distolingual cusp on upper molars. It differs from the proteo- terrestrial and aquatic vertebrate remains, including fishes, cro- pithecids by its less convergent orbits, P3–P4 with less developed codiles, and turtles20. The new primate from the PK2 locality of protocones and lacking incipient hypocone, upper molars with Pondaung Formation (Myanmar) does not belong to either of the more bunodont cusps, less developed buccal cingula, lacking two Asian anthropoid clades, the eosimiiforms1,3,6–8 and the hypocone and paraconule, weaker hypoparacrista, and stronger amphipithecids10–14, but shares instead derived dental characters lower canine. It differs from parapithecoids by its shorter muzzle, with some groups of the Eocene crown anthropoids and their less convergent orbits, less bunodont cusps on upper molars nearest fossil relatives. It therefore may bridge the gap between without additional enamel cusplets, unicuspid P2,P3–P4 length/ 3 3 Asian and African anthropoids. width proportions, and the absence of hypocone on P –M ,P2 as large as P3. It differs from the oligopithecids and the propliopithecids by its less convergent orbits, the retention of 2 3 4 Results P /2, absence of hypocone and hypoparacrista, P -P more Systematic paleontology. Order Primates Linnaeus, 1758 buccolingually extended, P3 with a more mesially located Suborder Anthropoidea Mivart, 1864 paracone and a sharper and longer postparacrista, with smaller Family incertae sedis protocone and no lingual cingulum, simple bulbous P3 with no Aseanpithecus gen. nov. honing facet and stronger lingual cingulid, nonconcave distal Aseanpithecus myanmarensis sp. nov. margin of their upper molars (except Catopithecus). It differs Etymology ASEAN, after the political and economic Union of from platyrrhines by its less convergent orbits, its smaller P2 and Southeast Asian countries and “pithecus” meaning monkey in more bunodont cusps on upper molars. It differs from Latin. The species name refers to the country where the fossil was Perupithecus by its larger molar size, more rectangular outline, discovered. larger trigon basin, weaker hypoparacrista, more bunodont cusps, Holotype NMMP 93 (Fig. 1a, b and Supplementary Fig. 1) left absence of metastyle, and more lingually located parastyle. maxilla with anterior orbital rim and C-M3 lacking M1 and buccal part of P4. Holotype and hypodigm fossils are housed in the Paleontology Collections of the National Museum, Ministry of Description. NMMP 93 (Fig. 1 and Supplementary Fig. 1): The Culture in Naypyitaw, Myanmar. left maxilla is nearly complete. It displays the alveolar region with Hypodigm NMMP 95 (Fig. 1c–g and Supplementary Fig. 1) C-M3 (buccal part of P4 and M1 are missing), the frontal anterior fragment of a right lower jaw with partial canine ascending process and the anterior rim of the orbit. It shows a – alveolus, P2 P3, and P4 alveoli and M1 mesial root, and NMMP unique maxillary component of the ventral orbital rim between 96 (Fig. 1h, i and Supplementary Fig. 1) right M3. the jugal and the upper margin of the ascending process. Lacrimal Type locality PK2 locality, Paukkaung village, Myaing Town- bone, foramen, and the associated canal are not present on the ship, Central Myanmar. The holotype and the hypodigms have smooth external surface and must have been therefore located been collected in very close proximity. inside the orbits, a characteristic anthropoid character21. In lateral Age Late Middle Eocene Pondaung Formation. view, its outline indicates an abbreviated and elevated muzzle Measurements (see Table 1) characterized by strong canine
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